Sphenacodon | |
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Skeleton of Sphenacodon ferox in the Field Museum of Natural History | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Synapsida |
Family: | † Sphenacodontidae |
Subfamily: | † Sphenacodontinae |
Genus: | † Sphenacodon Marsh, 1878 |
Species | |
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Synonyms | |
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Sphenacodon (meaning "wedge point tooth") is an extinct genus of synapsid that lived from about 300 to about 280 million years ago (Ma) during the Late Carboniferous and Early Permian periods. Like the closely related Dimetrodon , Sphenacodon was a carnivorous member of the Eupelycosauria family Sphenacodontidae. However, Sphenacodon had a low crest along its back, formed from blade-like bones on its vertebrae (neural spines) instead of the tall dorsal sail found in Dimetrodon. Fossils of Sphenacodon are known from New Mexico and the Utah–Arizona border region in North America.
Researchers currently recognize two species: Sphenacodon ferox (the type species) and Sphenacodon ferocior. Sphenacodon ferocior can be up to 40% larger in overall size (at about 3 m [9.8 ft] long) compared to Sphenacodon ferox (at about 2 m [6.6 ft]). In addition, the dorsal spines in Sphenacodon ferocior are proportionately 45% taller than in Sphenacodon ferox. The recent discovery [1] of a nearly complete skull of Sphenacodon ferox has helped clarify other distinctions between the two species, including the number of teeth in certain parts of the jaws and the size of the indented notch between the maxillary and premaxillary bones in the upper jaw. The two species occur together in some formations, but Sphenacodon ferox apparently survived later into the Early Permian.
Sphenacodon and Dimetrodon typically have been found in different geographical areas that were separated by the ancient Hueco Seaway that penetrated equatorial Pangaea during the Early Permian and "covered much of southern New Mexico and parts of West Texas". [2] Sphenacodon is known from the west in New Mexico, Arizona, and Utah, and Dimetrodon is known mainly from the east in Texas and Oklahoma in more deltaic environments. However, the species Dimetrodon occidentalis is found in New Mexico. [3] [4] Each genus would have been an apex land predator in its region and likely preyed on amphibians, diadectids, and early synapsids and diapsids. [5] Sphenacodon appears to have died out before about 280 million years ago during the Wolfcampian. [1] The genus Dimetrodon survived until about 270 million years ago. Such large sphenacodontid predators were later replaced by therapsids, the group of synapsids that includes the direct ancestors of mammals. [6]
The skull of Sphenacodon is very similar to that of Dimetrodon. [7] It is narrow from side to side and vertically deep, with an indented notch at the front of the maxillary bone in the upper jaw. The upper and lower jaws are equipped with an array of powerful teeth, divided into sharp pointed "incisors" [precaniniforms], large stabbing "canines" [caniniforms], and smaller slicing back teeth [postcaniniforms]. The orbit is set high and far back with a single opening (temporal fenestra) behind and partly below the eye, a characteristic of synapsids.
Body proportions are also similar to Dimetrodon, with a very large head, short neck, robust trunk, relatively short front and hind limbs, and a tapering tail that makes up about half the animal's entire length. However, the tops of the neural spines along the back bone are strikingly different in each genus. In Dimetrodon, the neural spines develop into long, narrow, cylindrical projections that support a tall vertical dorsal sail that ends near the base of the tail. In Sphenacodon, the neural spines are enlarged but retain a flat-tipped, blade-like shape along the back and tail, and form a crest rather than a tall sail. (The sphenacodontid genus Ctenospondylus also has blade-like neural spines, but its dorsal crest is taller than in Sphenacodon, although not as tall as the sail in Dimetrodon.)
There is evidence for strong epaxial muscles along the base of the raised neural spines in both Sphenacodon and Dimetrodon, likely helping to stiffen and strengthen the backbone for walking and for lunging at prey by restricting side-to-side flexing motion. A recent study [8] of the structure of the neural spines on Sphenacodon confirms that the upper parts were not encased in a thick muscular hump and instead protruded above a layer of muscle to form a low dorsal crest. Finds of sphenacodontid specimens in which postmortem distortion of the body caused the dorsal spines to overlap suggests that the spines were not connected by hard or particularly tough tissue. The possible function of a low, skin-covered crest in Sphenacodon is debated. A thermoregulatory role seems unlikely, although the taller crest in Sphenacodon ferocior is allometrically larger than in S. ferox. Recent research has favored a display role for the tall sails in Dimetrodon and Edaphosaurus. [9]
Both Sphenacodon and Dimetrodon have been depicted with their short limbs splayed outward at 90 degrees from the body in a wide pushup position and with the tail (and even belly) dragging on the ground, similar to modern lizards and crocodiles. A sprawling stance is also typical for Sphenacodon and Dimetrodon skeletons as currently mounted in museums. However, trackways called Dimetropus ("Dimetrodon foot") that match the foot configuration of large sphenacodontids show animals walking with their limbs brought under the body for a narrow, semi-erect gait without tail or belly drag marks. Such clear evidence for a more efficient upright posture suggests that important details about the anatomy and locomotion of Sphenacodon and Dimetrodon may not be fully understood. [10] Some well preserved narrow Dimetropus tracks found in parts of the Prehistoric Trackways National Monument in New Mexico match the smaller size of Sphenacodon, a genus known from skeletal fossils in the state, but could also come from a small Dimetrodon.
The American paleontologist O. C. Marsh [11] named Sphenacodon (from Greek sphen "wedge" + ake "point" + odous (-odon) "tooth") in 1878, based on part of a lower jaw (dentary) bone found in the redbeds of northern New Mexico by fossil collector David Baldwin. In his very short description of the jaw, Marsh cited the back teeth as characteristic ("crowns are much compressed, and have very sharp cutting edges without crenulations") and assessed the animal as "about six feet in length, and carnivorous in habit," although the rest of the skeleton was not known. He did not provide an illustration of the specimen. Marsh gave the genus the Latin specific name ferox "fierce" and erected the new family Sphenacodontidae, placed under the primitive reptilian order "Rhynchocephala" (=Rhynchocephalia), then including nearly all groups of early reptiles in addition to the living tuatara.
Other paleontologists overlooked Marsh's brief mention of Sphenacodon for almost three decades. [12] In the meantime, the sail-backed Dimetrodon, named in 1878 by rival paleontologist Edward Drinker Cope, became a scientifically important genus, known from numerous fossils. Recognition of Sphenacodon as a low-spined carnivorous "pelycosaur" distinct from Dimetrodon came in the early 20th century with the discovery of more fossils in New Mexico. [7] The proposed taxa Elcabrosaurus baldwini Case, 1907 and Scoliomus Williston and Case, 1913 now are considered junior synonyms of Sphenacodon ferox.
In 1937, Alfred Sherwood Romer [13] described a second species from New Mexico named Sphenacodon ferocior ("fiercer") that was larger and more robust, with proportionately longer neural spines. Romer and Price (1940) [7] provided detailed descriptions of both ferox and ferocior with skeletal reconstructions. [14]
A third species, Sphenacodon britannicus, has sometimes been cited in the literature. In 1908 German paleontologist F. von Huene [15] described Oxyodon britannicus, based on part of a maxilla found in England, (The generic name Oxyodon is preoccupied by a fish (Oxyodon Baur, 1906) and so is invalid.) The specimen had been identified earlier as a possible Triassic dinosaur, but von Huene recognized a "pelycosaur." Paton [16] transferred the species to Sphenacodon in 1974, noting it would have been an animal about the size of Sphenacodon ferox. However, more recent studies [17] have questioned whether such limited fossil material can be used to distinguish between Dimetrodon and Sphenacodon—or its own genus. The species "Oxyodon" britannicus (or as Sphenacodon (?) britannicus) is now generally classified as Sphenacodontidae incertae sedis (of uncertain placement). [1] [8]
Sphenacodon in a cladogram after Fröbisch et al., 2011: [18]
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Dimetrodon is a genus of non-mammalian synapsid that lived during the Cisuralian age of the Early Permian period, around 295–272 million years ago. It is a member of the family Sphenacodontidae. With most species measuring 1.7–4.6 m (5.6–15.1 ft) long and weighing 28–250 kg (62–551 lb), the most prominent feature of Dimetrodon is the large neural spine sail on its back formed by elongated spines extending from the vertebrae. It was an obligate quadruped and had a tall, curved skull with large teeth of different sizes set along the jaws. Most fossils have been found in the Southwestern United States, the majority of these coming from a geological deposit called the Red Beds of Texas and Oklahoma. More recently, its fossils have also been found in Germany and over a dozen species have been named since the genus was first erected in 1878.
Pelycosaur is an older term for basal or primitive Late Paleozoic synapsids, excluding the therapsids and their descendants. Previously, the term mammal-like reptile had been used, and pelycosaur was considered an order, but this is now thought to be incorrect, and seen as outdated.
Edaphosaurus is a genus of extinct edaphosaurid synapsids that lived in what is now North America and Europe around 303.4 to 272.5 million years ago, during the Late Carboniferous to Early Permian. American paleontologist Edward Drinker Cope first described Edaphosaurus in 1882, naming it for the "dental pavement" on both the upper and lower jaws, from the Greek edaphos έδαφος and σαῦρος ("lizard").
Sphenacodontidae is an extinct family of sphenacodontoid synapsids. Small to large, advanced, carnivorous, Late Pennsylvanian to middle Permian "pelycosaurs". The most recent one, Dimetrodon angelensis, is from the latest Kungurian or, more likely, early Roadian San Angelo Formation. However, given the notorious incompleteness of the fossil record, a recent study concluded that the Sphenacodontidae may have become extinct as recently as the early Capitanian. Primitive forms were generally small, but during the later part of the early Permian these animals grew progressively larger, to become the top predators of terrestrial environments. Sphenacodontid fossils are so far known only from North America and Europe.
Sphenacodontia is a stem-based clade of derived synapsids. It was defined by Amson and Laurin (2011) as "the largest clade that includes Haptodus baylei, Haptodus garnettensis and Sphenacodon ferox, but not Edaphosaurus pogonias". They first appear during the Late Pennsylvanian epoch. From the end of the Carboniferous to the end of the Permian, most of them remained large, with only some secondarily becoming small in size.
Eupelycosauria is a large clade of animals characterized by the unique shape of their skull, encompassing all mammals and their closest extinct relatives. They first appeared 308 million years ago during the Early Pennsylvanian epoch, with the fossils of Echinerpeton and perhaps an even earlier genus, Protoclepsydrops, representing just one of the many stages in the evolution of mammals, in contrast to their earlier amniote ancestors.
Dimetrodon borealis, formerly known as Bathygnathus borealis, is an extinct species of pelycosaur-grade synapsid that lived about 270 million years ago (Ma) in the Early Middle Permian. A partial maxilla or upper jaw bone from Prince Edward Island in Canada is the only known fossil of Bathygnathus. The maxilla was discovered around 1845 during the course of a well excavation in Spring Brook in the New London area and its significance was recognized by geologists John William Dawson and Joseph Leidy. It was originally described by Leidy in 1854 as the lower jaw of a dinosaur, making it the first purported dinosaur to have been found in Canada, and the second to have been found in all of North America. The bone was later identified as that of a pelycosaur. Although its current classification as a sphenacodontid synapsid was not recognized until after the discovery of its more famous relative Dimetrodon in the 1870s, Bathygnathus is notable for being the first discovered sphenacodontid. A 2015 study by the researchers from the University of Toronto Mississauga, Carleton University and the Royal Ontario Museum reclassified the species into the genus Dimetrodon.
Neosaurus is an extinct genus of pelycosaur-grade synapsids from the Late Carboniferous-Early Permian of the Jura region of France. It is known only from a partial maxilla or upper jaw bone and an associated impression of the bone. The teardrop shape of the teeth in the jaw indicate that Neosaurus belongs to the family Sphenacodontidae, which includes the better-known Dimetrodon from the Southwestern United States. The maxilla was first attributed to an early diapsid reptile in 1857, and later a crocodylomorph in 1869, before finally being identified as a sphenacodont synapsid in 1899, a classification that still holds today.
Varanopidae is an extinct family of amniotes that resembled monitor lizards and may have filled a similar niche, hence the name. Typically, they are considered synapsids that evolved from an Archaeothyris-like synapsid in the Late Carboniferous. However, some recent studies have recovered them being taxonomically closer to diapsid reptiles. A varanopid from the latest Middle Permian Pristerognathus Assemblage Zone is the youngest known varanopid and the last member of the "pelycosaur" group of synapsids.
Ctenospondylus is an extinct genus of sphenacodontid synapsid
Secodontosaurus is an extinct genus of "pelycosaur" synapsids that lived from between about 285 to 272 million years ago during the Early Permian. Like the well known Dimetrodon, Secodontosaurus is a carnivorous member of the Eupelycosauria family Sphenacodontidae and has a similar tall dorsal sail. However, its skull is long, low, and narrow, with slender jaws that have teeth that are very similar in size and shape—unlike the shorter, deep skull of Dimetrodon, which has large, prominent canine-like teeth in front and smaller slicing teeth further back in its jaws. Its unusual long, narrow jaws suggest that Secodontosaurus may have been specialized for catching fish or for hunting prey that lived or hid in burrows or crevices. Although no complete skeletons are currently known, Secodontosaurus likely ranged from about 2 to 2.7 metres (7–9 ft) in length, weighing up to 110 kilograms (250 lb).
Mycterosaurus is an extinct genus of synapsids belonging to the family Varanopidae. It is classified in the varanopid subfamily Mycterosaurinae. Mycterosaurus is the most primitive member of its family, existing from 290.1 to 272.5 MYA, known to Texas and Oklahoma. It lacks some features that its advanced relatives have.
Eothyris is a genus of extinct synapsid in the family Eothyrididae from the early Permian. It was a carnivorous insectivorous animal, closely related to Oedaleops. Only the skull of Eothyris, first described in 1937, is known. It had a 6-centimetre-long (2.4-inch) skull, and its total estimated length was 30 centimetres. Eothyris is one of the most primitive synapsids known and is probably very similar to the common ancestor of all synapsids in many respects. The only known specimen of Eothyris was collected from the Artinskian-lower.
Ctenorhachis is an extinct genus of the family Sphenacodontidae. Ctenorhachis lived in the Early Permian epoch. Ctenorhachis was related to Dimetrodon, but did not belong to the same subfamily as Dimetrodon and Sphenacodon, being a more basal member of Sphenacodontidae. Two specimens are known that have been found from the Wichita Group outcropping in Baylor and Archer counties, north-central Texas. Only the vertebrae and pelvis are known. Articulated vertebrae from the holotype specimen possess blade like neural spines that are greatly enlarged, although not nearly to the extent that can be seen in more derived sphenacodontids such as Dimetrodon and Secodontosaurus, in which they form a large sail. The pelvis is nearly identical to that of Dimetrodon. As suggested in the original description of the genus, Ctenorhachis may represent a short-spined sexual dimorph, although the authors find this unlikely.
Aerosaurus is an extinct genus within Varanopidae, a family of non-mammalian synapsids. It lived between 252-299 million years ago during the Early Permian in North America. The name comes from Latin aes (aeris) “copper” and Greek sauros “lizard,” for El Cobre Canyon in northern New Mexico, where the type fossil was found and the site of former copper mines. Aerosaurus was a small to medium-bodied carnivorous synapsid characterized by its recurved teeth, triangular lateral temporal fenestra, and extended teeth row. Two species are recognized: A. greenleeorum (1937) and A. wellesi (1981).
Eosyodon is a dubious genus of extinct non-mammalian synapsids from the Permian of Texas. Its type and only species is Eosyodon hudsoni. Though it was originally interpreted as an early therapsid, it is probably a member of Sphenacodontidae, the family of synapsids that includes Dimetrodon.
Echinerpeton is an extinct genus of synapsid, including the single species Echinerpeton intermedium from the Late Carboniferous of Nova Scotia, Canada. The name means 'spiny lizard' (Greek). Along with its contemporary Archaeothyris, Echinerpeton is the oldest known synapsid, having lived around 308 million years ago. It is known from six small, fragmentary fossils, which were found in an outcrop of the Morien Group near the town of Florence. The most complete specimen preserves articulated vertebrae with high neural spines, indicating that Echinerpeton was a sail-backed synapsid like the better known Dimetrodon, Sphenacodon, and Edaphosaurus. However, the relationship of Echinerpeton to these other forms is unclear, and its phylogenetic placement among basal synapsids remains uncertain.
Macromerion is an extinct genus of non-mammalian synapsids, specifically Pelycosaurs, in the family Sphenacodontidae from Late Carboniferous deposits in the Czech Republic. It was named as a species of Labyrinthodon in 1875 and as its own genus in 1879.
Gordodon is an extinct genus of non-mammalian synapsid that lived during the Early Permian of what is now Otero County, New Mexico. It was a member of the herbivorous sail-backed family Edaphosauridae and contains only a single species, the type species G. kraineri. Gordodon is unusual among early synapsids for its teeth, which were arranged similarly to those of modern mammals and unlike the simple, uniform lizard-like teeth of other early herbivorous synapsids. Gordodon had large incisor-like teeth at the front, followed by a prominent gap between them and a short row of peg-like teeth at the back. Gordodon was also relatively long-necked for an early synapsid, with elongated and gracile vertebrae in its neck and back. Like other edaphosaurids, Gordodon had a tall sail on its back made from the bony neural spines of its vertebrae. The spines also had bony knobs on them, a common trait of edaphosaurids, but the knobs of Gordodon are also unique for being more slender, thorn-like and randomly arranged along the spines. It is estimated to have been rather small at 1 m in length excluding the tail and 34 kg (75 lb) in weight.